Ibeas (Spain)

J. L. Arsuaga, J. M. Carretero, I. Martinez & A. Gracia

Cranial remains and long bones from Atapuerca/lbeas (Spain)

Dpto. de Paleontologfa. Fat. de CC. Geoldgicas, Universidad Comphtense, Z&W Madrid, Spain

The cranial remains and long bones of the Sima de 10s Huesos site (Sierra de Atapuerca, Spain) are described and their phylogenetic affinities established. We find in the Atapuerca/Ibeas sample: (a) a number ofplesiomorphic traits (namely, character states not retained by the upper Pleistocene Neandertals but present in the outgroups); (b) one apomorphy shared with Neandertals; (c) some postcranial character states shared with Neandertals but whose phylogenetic status remain uncertain. In sum, the Atapuerca/Ibeas fossils are phylogenetically related to the Neandertals but cannot be pooled with them because of the overall plesiomorphic pattern. Other European middle Pleistocene specimens show apomorphies shared with Neandertals and plesiomorphies. In our opinion these specimens do not share uniquely derived features and thus this chronological group cannot be defined. Other aspects such as cerebral lateralization and sexual dimorphism in humeri and temporal bones are investigated.

Received 20 July I989 Revision received 15 January 1990 and accepted 15 June 1990 ITeyroords:Atapuerca, middle Pleistocene, Neandertals, cranial remains, long bones.

Journal of Human Evolution ( 199 1) 20,19

The Atapuerca

l-230

sites

The Sierra de Atapuerca (Burgos, Spain) consists of a monadnock of Cretaceous limestones, to the west ofand detached from the main ranges of the Iberic System. Several fossil-bearing cave fillings are known in the wide karst network of the Atapuerca

Hill (Figure

1). Some of

them were exposed by a railway trench and have been excavated or sampled since 1976. These deposits are bracketed within the limits of the middle Pleistocene. Most attention has been paid to the TD and TG sites where important assemblages have been found. A large number fossil-bearing

of human

remains

archaeological

and palaeontological

(more than 200) have been recovered

from another

cave filling, Sima de 10s Huesos (SH), less than 1 km from TD-TG

ing to another karst system, Cueva Mayor-Cueva The SH deposit lays in a small blind ch,amber

but belong-

de1 Silo, in the Ibeas de Juarros near the bottom

township.

of a 13 m deep pit, more

than 500 m from the Cueva Mayor current entrance. Unfortunately, many tons of bone bearing breccia were removed before 1976 by excursionists in search of bear canines and bones, and left aside in the same room as a mass ofbone fragments and mud debris. In 1976, 1983 and following years some 5 tons ofoverburden material has been excavated, washed and scrutinized. The fossil assemblage Neither herbivores with a minimum

of the Sima de 10s Huesos debris consists of carnivores

number

of individuals

based on some 9000 fossils, is unequivocal

close to 200. The identification (Torres,

of the human fossils has been dated by gamma-ray (U-Th

and humans.

nor stone tools have been found. The taxon Ursus deningeri is dominant,

age) and > 175 ky (U-Pa age) (Yokoyama,

of Ursus deningeri,

1978, 1984, 1987; Torres et al., 1978). One spectrometry

to 320 ky + 233 ky/-73 ky

1989).

The human fossils The minimum number of individuals has been estimated as 11. Some preliminary or partial studies on the Atapuerca/Ibeas cranial remains and long bones have been published by Martinez & Arsuaga (1985, 1987), P Crez & Bermlidez de Castro (1985), Perez ( 1987) and 0047-2484/91/030191+40

$03.00/O

0 1991 Academic

Press Limited

J.

192

et al. (1989).

Aguirre identified

Up to 1988, 96 human

and labelled.

we will analyse

Some

of them

the most complete

1. Frontal bone remains in 1986). This

AT-121 (found

medially

ing to the terminology general

good

severely

consists

damaged break.

exposed

diploe.

surface

structure

(19806),

aspect

is greater

than

Wolpoffs

results,

European

Western

middle ofAT-

in Petralona the data

surface

aspect

in the orbital consists

corner

of the bone

and

was produced

of this section,

tables

by the

and

internal

two small cavities

sinus extends

well laterally

of the

into the

1973, 1978, 1979, 1980, 1983) is in the vicinity

have

trigone.

of the temporal

and

(1982)

the same

for Arago

According

supraorbital

but the supraorbital

Steinheim

of Spitery

supraorbital

Steinheim

is in

but it is

portion).

of the bone

(Tappen,

of

(accord-

of the torus,

parasagittal)

of the torus, especially

specimen, and

Neandertals

and

segment

in this fragment

where the external surface is preserved. torus in AT- 12 1 can be described as projected,

Petralona

than for any Neandertal

superior

the frontal

and with a large and convex

Bilzingsleben,

and

of the torus

line, as well as on the anterior aspect The preserved part of supraorbital

greater

been

torus

So, the lateral

1980). The external

torus. configuration

in the superior

developed

have

In this paper

of a left supraorbital

is missing.

(not exactly

in AT-121

of the supraorbital pattern ofsurface

distinctive

fragments

1).

are represented

(especially

of the torus

Thus,

notch

squama

In the postero-inferior

sinus are exposed.

fossils (Table

segment

segment

& Ranyard,

in the frontal

in the anterior

The

orbital segment A vermiculate

evenly

and can be associated.

of a large

the orbital

of Smith

cross section

coarse cancellous

clearly

and long bone

temporal line, orbital plate and temporal fossa, as well as an suture (Figure 2). The supraorbital torus extends from this

halfof

condition

A triangular

frontal

cranial conjoin

for 54 mm, but the supraorbital

the torus and the lateral

medial

clearly

and representative

surrounding portions ofsquama, almost complete frontozygomatic suture

ET AL.

L. ARSUAGA

midorbit

torus length

21, Steinheim,

thickness

of La Quina

5

Contrary

to

as Bilzingsleben.

do not show any clear difference

thick,

to Wolpoff

Petralona

in thickness

and

between

six the

Pleistocene and the Neandertal specimens. We have taken the supraorbital thickness (after Smith & Ranyard, 1980) at the midorbit and lateral points (equivalent to

the minimum thickness and the supraorbital trigone thickness of Spitery, 1982). The values obtained (Table 2) are near or beyond the upper limit of the sample ranges of Spitery ( 1982) (European

( 1980) defined

middle

(European by Smith

Pleistocene

plus western

Neandertal

South-Central Neandertals). & Ranyard, 1980) of AT-121

specimens)

The supraorbital are greater than

South-Central Neandertal ranges and similar to Arago 2 1. The frontozygomatic suture is preserved in AT-121, except constituting

a very extensive

surface

(19 mm x 11 mm).

and Smith

in its most

The distance

& Ranyard

torus projections (as the upper limits of the

fmofmt

posterior

apex,

can be esti-

mated to be 11 mm. According to the figures of Bouzat (1982), the range for a sample of four Western Neandertals and three European middle Pleistocene fossils (Arago 21, Petralona, Steinheim) is 9-l 3 mm. The orbital plate of AT-121 is preserved 37 mm x 26 mm) bounded by the supraorbital matic suture and, posteriorly, a line of fracture internal table of the frontal bone and the orbital Above preserved

in a rectangular area (of approximately margin, the medial break, the frontozygocorresponding to the junction between the plate.

the torus, and separated from it by the supratoral sulcus, the frontal squama is to a maximum length of 41 mm. The squama thickness is 9-5 mm at the most

CRANIAL

REMAINS

AND

LONG

BONES

FROM

193

ATAPUERCA/IBEAS

N (u.T.M.)

Cueva del

Corn

G. del Silex

SH ueaos

4&

Grupo Espoleal6gico Edelweiss (1977-79) Ercma. Diputacih Provincial de Burgas

Figure 1. The karst network of the Atapuerca Hill. Arrows point to the Cueva Mayor and Cueva de1 Silo entrances. Asterisks mark the fossil-bearing cave fillings which have been excavatedor sampled.Courtesy of the Grupo EspeleblogicoEdelweiss(Excma. Diputaci6nProvincialde Burgos).

posterior preserved point. The temporal line is superficially

altered but persists in strong relief

above the deep temporal notch. The frontal squama seems to be considerably to a horizontal plane defined by the orbital roof. The endocranial surface of AT- 12 1 is preserved

angled relative

as far as 30 mm up from the junction

orbital roof/internal table ofthe frontal bone. Most of this surface is clearly depressed in a sort ofdigitation or fossa which seems to correspond to the middle frontal convolution. AT-200

(found in 1988).

This frontal fragment

includes the medial half of a right supra-

orbital torus and a great portion of the interorbital region (Figure 2). Almost all the preserved exocranial surface belongs to the right side of the frontal bone, the supraorbital torus

194

J. L. ARSUAGA

AtapuercajIbeas

Table 1

human fossils

ET AL.

studied in this article

Partial skulls -Cranium 1: Composed of Parietal III + temporal fragment AT-86f occipital fragment AT-122. --Cranium 2: Composed of frontal fragment AT-36 + Parietal II + wormian bone AT-76 +occipital fragment AT-66. Isolated cranial bones -Frontal bone: AT-121, AT-200, AT-174, AT-129, AT-50/AT-52. -Parietal bone: Parietal I. -Occipital bone: Occipital I, Occipital II, Occipital III, AT-122, AT-39, AT-123a/AT-123b. -Temporal bone: AT-84, AT- 124, AT-220, AT- 125. Long bones -Humerus: AT-25, AT-93, AT-2 17. -Ulna: AT-2 18. -Tibia: Tibia I, AT-85, AT-IS.

extending

from the glabella

supraorbital

laterally

torus. The endocranial

for 55 mm. The orbital roof is preserved beneath surface ofAT-

the

extends on the right side 35 mm from

the frontal crest and 16 mm on the left side. Only the beginning of the frontal squama, above the glabellar

region, is present, extending

as far as 33 mm above the nasofrontal

suture.

The frontal fragment is broken on the left side short ofthe midline. The cross section reveals a well developed frontal sinus which extends medially reaching the midsagittal plane. The dimensions of the preserved chamber are: height = 20 mm, width = 16 mm, antero-posterior depth = 10 mm. The sinus is limited posteriorly by the internal table of the frontal bone and anteriorly

by a well developed sinus wall (of some 10 mm) made of external

bone table and

(mostly) coarse cancellous bone. This sinus cavity does not enter into the frontal squama and its lateral extension is unknown. On the right side the supraorbital torus is broken in a very oblique section (from lateral to medial). This cross section does not exhibit any frontal sinus, showing an internal

structure

of coarse cancellous

tables. The right side of the nasofrontal

bone between the internal

suture is totally preserved

and external

as well as part of the

frontomaxillary suture. An inferior break exposes on the right side another frontal sinus chamber which extends upwards much less that the left one (dimensions = 13 x 13 x 10 mm). Thus, there is a clear asymmetry

in frontal pneumatization

in AT-200.

Limited by these two

big frontal sinuses, the nasofrontal suture and the frontal crest, there is also a small and poorly developed sinus cavity (6 x 7 x 10 mm). According to Tillier ( 1977: 287) “constant pneumatization of the supraorbital torus is a characteristic feature of Western European Neandertal men and their predecessors of the Riss-Wiirm interglacial”. Nevertheless, we do not believe that frontal sinus development European

can be used to establish

phylogentic

affinities,

at least in

human evolution.

The external surface is in general good condition, with some small patches of superficial alteration. There are many vascular foramina in the surface (as in AT- 12 1) . A clear vermiculate pattern is present on the anterior and superior surfaces of the torus in the segment lateral to the supraorbital notch, which is medially substituted by a less convoluted pattern. In the glabellar segment the surface shows a pattern of grooves and small pits. In the superior orbital margin there is a well marked supraorbital notch and 6 mm medial to it a narrow groove can be observed: the internal frontal notch. There is neither a supraorbital foramen above the notch nor a supraorbital tubercle. In AT-200 there is no

CRANIAL

REMAINS

AND

LONG

BONES

FROM

ATAPUERCAIIBEAS

Figure 2. Frontal bones. (1) AT-200: Anterior view; (2) AT-200: Left transversal break; (3) AT-200: Right transvcrsc break; (4) AT-200: I IIfprior view; (5) AT-121: Anteriorview; (6) AT-121: Lateralview; (7) ATI2 I : Medial transverse break; (81 AT- I2 I : Superior view.

195

196

J. L. ARSUAGA

ETAL.

Projection Sample

Krapina Average S.D. x Range Sala

Lateral

24.3 1.4

a

23@ 27.0 25.0

Central and eastern European Early Upper Palaeolithic Average 20.3 S.D. 2.6 N 9 Range 15.G 23.0 La Quina 5 21 La Chapelle-auxSaints Arago 2 1 (on cast) 34 Steinheim (on cast) AT-121 35 AT-200

Midorbit

Thickness Medial

Lateral

Midorbit

Medial

23.9 1.2 11 23.026.0 22.5

20.3 2.3 4 17.523.0 20.0

12.5 1.6 11 10.316.0 11.0

IO.7 1.8 13 7.014.3 7.1

17.6 3.0 4 15.822.0 15.0

Wolpoff (1981)

16.1 3.4 9 8+ 19.0 21

13.0 3.0 9 8.0 17.5 22

8.1 1.4 11 6.@ IO.1 9.5

5.4 1.7 11 447.7 9

16.6 3.3 11 11.523.7 13

Wolpoff etal. (1981)

11.5 11.5 10 14 -

10 10 10 14

13.5 16

29

20

31 (26)’

;:6)?

( 16.5)3

source

et

al.

Wolpoff et al. (1981)

Authors’ Authors’ Authors’ Authors Authors Authors

‘Averages between both sides. ‘This figure must be considered as a minimum value taken in the most medial point of the preserved “This figure was taken medial to the standard point.

torus.

supraorbital sulcus extending from the supraorbital notch, so this fossil exhibits a true supraorbital torus with complete fusion of the superciliary arch and the supraorbital arch. The glabellar region has an inflated appearance and no glabellar depression can be seen in the midline. The superciliary arches and the glabellar are so continuous that the arches cannot be isolated and there is a complete fusion between the supraorbital and glabellar tori in an uninterrupted, continuous bone shelf. In AT-200 and Bilzingsleben the glabellar torus forms a very large bulge of similar dimensions. The distance from nasion to frontal crest is 25 mm in both Bilzingsleben and AT200, and from glabella to frontal crest 28 mm in Bilzingsleben and 30 mm in AT-200. However, it is better to avoid the frontal crest when measuring the glabellar projection; then we obtain 25 mm in AT-200 and only 18 mm in La Quina 5. In AT-200 the torus emerges very gradually from the frontal squama, so that there is not a well-marked supraglabellar sulcus but only a very small shallow supraglabellar fossa in the midline. Bilzingsleben is also described as lacking a true supraglabellar sulcus (ViEek, 1978). The interorbital region of AT-200 is very wide and there is no nasal root depression. In fossil and modern humans the frontal’s margo nasalis shows an inverted V shape in anterior view, but in AT-200 the medial segment of the nasofrontal suture follows a horizontal course. This atypical articulation between frontal and nasal bones can also be found in Arago 2 1 and “Sinanthropus” skull XII. The nasofrontal suture is very thick in AT-200 (14 mm at the nasion).

CRANIAL

Though

thickness

REMAINS

197

AND LONG BONES FROM ATAPUERCA/IBEAS

and projection

of the supraorbital

standard points, AT-200 appears, in the preserved specially very projecting (Table 2).

torus cannot

supraorbital

be measured

segment,

at the

very thick and

The internal surface of the frontal bone is in good condition. Only the frontal crest, preserved for a length of 14 mm, shows some erosion. On the right side impressions of the first and second frontal convolutions

can be found as well as that ofthe first frontal convolution

on

the left side. The right frontal first convolution is more developed than the left one, so a right frontal petalia is present in terms of the more anteriorly projecting (4.5 mm) right frontal pole. This petalia suggests right-handedness.

Bermlidez

buccal striations on the anterior teeth of the Atapuerca

de Castro et al. (1988) have studied dental sample and some Neandertals.

These authors support the hypothesis that the striations were produced when stone tools were used to cut something

held between

the anterior

teeth. The pattern

of striations

indicates

right- or left-handedness. These results provide indirect evidence for lateralization of the brain in fossil humans. In the Atapuerca sample buccal scratches were detected on 23 anterior teeth belonging to at least eight individuals (BermGdez de Castro, personal communication).

In all cases right-handedness

is concluded.

AT-174 (found in 1986). This fossil represents a small portion of a right supraorbital the orbital segment),

torus (in

preserving parts of the superior aspect ofthe torus, anterior aspect (very

eroded), orbital plate and endocranial surface. The maximum diameter of the fragment is 34 mm. The two surfaces of fracture that limit transversally the torus show triangular cross sections similar, in size and internal structure, to that of AT-121. At the medial break two alveolar concavities are exposed marking the most lateral extent of the frontal sinus. AT-129

(found in 1986).

vermiculate maximum AT-50 squama

surface

This is a triangular

pattern

exocranially.

fragment

Its

of frontal

maximum

diameter

thickness is 13 mm. There is no sutural preservation

and AT-52

(recovered

that clearly match

in 1984). These are two fragments

(maximum

diameter

squama

exhibiting

is 39 mm

and

a the

on its edges. (lacking sutures) offrontal

of the two pieces combined = 52 mm). No

vermiculate surface pattern is present on the external surface. AT-52 shows, on the endocranial surface, a short segment ( 14 mm long) of the sagittal sulcus, appearing as a shallow groove enclosed between two smooth edges of the frontal crest. The thickness of these remains is very constant and varies from 8 to 10 mm. AT-36

(recovered

in 1984). A rectangular

(24 mm). It is very constant Parietal

in thickness

(43

x

26 mm) with a segment of coronal

(maximum

8.5 mm). This piece articulates

suture with

II along the coronal suture.

2. Parietal bone remains Parietal I (P. I). This is not a single fragment only one specimen and recovered

but a set of five parietal pieces belonging

to

in three different years: AT-l 7 (found in 1976), AT-3la/

AT-31b (found in 1984) and AT-173a/173b (retrieved in 1987). Once the five fragments are combined, most of a left parietal bone is represented (Figure 3). The parietomastoid suture is complete but the edge is abraded, lacking indentation. Its length (distance from the aster-ion to the parietal notch) is about 33 mm, a value similar to those of Arago 47 (32 mm) and Swanscombe parietal left (28 mm). Parietal I preserves also segments of the lambdoidal (27 mm) and coronal (11 mm) borders, both showing denticulation. The squamous suture, or the area just above it, is preserved as far as 28 mm anterior

to the parietal notch. Some

198

J. L. ARSUAGA

ET AL.

Figure 3. Parietal bones. (I) C ranium 2: Superior view. This specimen is made up of Parka1 11 plus the frontal fragment AT-36 and the wormian (lambdatic) bone AT-76, which connects with the occipital fragment AT-66; (2) Parietal I; (3) Parietal III with the occipital fragment AT-122. They comprise. together with the temporal fragment AT-86, the Cranium 1.

CRANIAL

Table 3

199

REMAINS AND LONG BONES FROM ATAPUERCA/IBEAS

Parietal thickness (mm)

European Middle Pleistocene La Chaise (Suard) Steinheim Arago 47 Swanscombe Caste1 di Guido Fontechevade Biache-Saint-Vaast Atapuerca P.1 Atapuerca P.11 Atapuerca P.111 Neandertal Parameters Average S.D. N Range “Sinandropus” Parameters Average S.D. .N Range

Parietal boss

1

9

6.5 8.5 11.5 8 6 13 z-102

Asterion

Source

9.5 (11.5)3

Piveteau (1970) Wolpoff (19806) Authors (on cast) Wolpoff ( 19806) Mallegni et al. (1983) Vallois ( 1958) Vandermeersch (1982) Authors Authors Authors

7.1

Wolpoff (19806)”

8 6.5 14.5 10.1 12.5 8 8 (9)’

8.3 1.69 12 5-l 1.3

10 5-8.9

11.8 2.1 6 9-16

14.8 1.5 8 13.5-I 7.4

1.68

Weidenreich

( 1943)5

‘8 mm at the suture and 9 mm at the superior temporal line level near the asterion. *As the p&eta1 boss is missing the thickness was taken in its vicinity, so that the real value is probably greater. 99.5 mm at the lambdoidal suture and 11.5at the point where the superior temporal line crosses the suture near the asterion. *Calculated by the authors on Wolpofl’s measures at middle eminence and mastoid angle for Central and Western

European Neandertals. ‘Calculated by the authors on Weidenreich’smeasures. ridges, related to the articulation (overlap) between the temporal squama and the parietal bone, are present. The external surface is preserved in good condition except for two small areas of outer table loss in AT-3 1a and AT- 17. Parietal I is very thick, attaining a thickness value of 13 mm at the parietal protuberance. This value is higher than for any European middle Pleistocene or Neandertal specimen in Table 3, and only one “Sinunthropus” skull has a greater thickness at the parietal boss. The temporal lines are visible on the external surface but they are only slightly marked in their posterior part. The superior temporal line does not finish in an angular torus at the mastoid angle. For this reason the Parietal I thickness at asterion (8 mm in the suture and 9 mm at the level of the superior temporal line near the asterion) is clearly smaller than for Arago 47 and the “Sinanthropus” specimens, which show conspicuous angular tori (Table 3). This structure has been described on “Sinanthropus” (Weidenreich, 1943), Dali (Wolpoff et al., 1984) and several Javanese specimens (from Sangiran, Sambungmachan and Ngandong: Grimaud, 1982a, 1986) as well as on the African fossils from Bodo (Asfaw, 1983)) Broken Hill and OH 9. Mallegni et al. (1983) have reported the presence ofangular torus on Caste1 di Guido 5, but in our opinion (based on a cast) the mastoid angle is thick but without a well defined torus. To Grimaud (19826) Petralona exhibits angular torus, but to Stringer (1984) the trait is absent in this fossil. In sum, only Arago 47 in the European fossil record exhibits an irrefutable angular torus.

200

J. L. ARSUACA

ET AL.

Parietal I lacks the sagittal suture but the curvatures of the complete biparietal vault can be tentatively interpreted with the help of P. II. Parietal I shows a regular curvature, reaching its maximum

(parietal

superior temporal

line. This point would correspond

protuberance)

in the middle of the bone, slightly

parietal walls would appear slightly convergent Arago 47, Petralona

or Swanscombe.

above the

to the parietum. In norma occipitalis the

toward the top (or nearly parallel)

In our opinion,

the neandertal-derived

pattern

as in “en

bombe” was not present in P. I. There

is a pathological

erosion near the postero-superior

damage is oval sized (maximum

corner

diameter = 25 mm), well delimited

of P. I (AT-l 7). The and only affecting

the

external bone table without diploe exposition. There are signs of bone regeneration. In the deepest zone of this lesion the surface is rough. The most likely etiology would be a traumatic injury produced by a blunt object, followed by a very restricted infectious disturbance (P. J. PCrez, personal communication). Parietal I exhibits well marked vascular impressions on its endocranial surface (Figure 4). A high anastomotic

degree is observed,

especially

network that covers the whole endocranial lateral sinus. In modern populations

the impression

of a slender

surface. Neither Atapuerca

capillary

P. I nor P. III display

this sulcus runs from the occipital to the temporal bone,

usually crossing the asterionic region of the parietal bone, where a deep groove can be easily identified at this point. In the Atapuerca temporal bones AT-84 and AT-124 the sulcus courses from the occipital

to the temporal

present in the early middle Pleistocene

bone below asterion.

A clear lateral

parietal bone from Ternifine

(Arambourg,

sulcus is 1963) but

in the “Sinanthropus” Weidenreich (1943).

crania the sulcus does not extend to the parietal bone according to In the European middle Pleistocene the sulcus is either completely

absent or represented

only by the superior edge of the groove. A lateral sulcus on the parietal

bone is said to be common

in Neandertals

but it is lacking in Cova Negra (Arsuaga

et al.,

1989) and not clearly represented in La Quina 5, Krapina 3 (cast) and 6 (cast). The fragment AT-17, corresponding to the superior plane of P. I, was oriented upside down by Saban (1980, 1982, 1984, 1985, 1986) and considered temporal plane. So, the P. I vascular impressions were wrongly described and misunderstood by this author. A full description of the meningeal

vascular

system and cerebral

impressions

in the Atapuerca

sample will be

published elsewhere.

Parietal

II (P. II). Five left parietal fragments

have been combined

in Parietal

II: AT-18

(recovered in the 1976 season), AT-33 and AT-61 (both found in 1984), AT-1 26 (found in 1986) and AT-2 11 (found in 1988). In the following paragraphs we will consider all the five fossils as a single specimen, representing a great part of a left parietal postero-inferior quadrant, the parietal protuberance and a rectangular

bone, lacking the portion (of some

42 x 25 mm) containing the bregma and the adjacent segments of the coronal and sagittal sutures. Parietal II connects with the frontal fragment AT-36 and with the wormian bone AT-76,

which matches with the occipital fragment AT-66,

Cranium 2 (Figure 3). Parietal II preserves 34 mm ofcoronal

the complete set being assigned to

border (all the suture lengths are given as chords in

this paper) in the fragment AT-61 and 9 mm in AT-2 11. The sagittal suture is represented by two disjointed segments (28 mm in AT-33 and 19 mm in AT-l 8), as well as the lambda (in AT- 18) and a portion of the lambdoidal border, which can be divided into two segments (of 25 mm each) showing strong angulation (130”). Th is atypical course of the lambdoidal suture is due to the existence ofextra-sutural bones. One of them, AT-76 (found in 1984), is

CRANIAL

REMAINS AND LONG BONES FROM ATAPUERCAIIBEAS

-. -. -. =.. .Z

--....__ -. -.

: ; : : : .. .: :. : :. . :. : : : :. :. . : : :

-.

201

202

J. L. ARSUAGA

entirely

preserved

articulates

with

fragment

AT-66.

with

all borders.

Parietal

II

Roughly

(for 23 mm

According

ET

rhomboidal

from

to Wolpoff

AL.

in shape

the lambda),

(1980a),

(32 x 34 mm),

as well as with

Vertesszollos

and Petralona

AT-76

the occipital also have large

extra-sutural bones at the top of the occipital bone (but to Thoma, 1978, the possibility of an anomalous bone in the lambda region of Vertesszollos is very remote). To Wolpoff ( 19800: 230) “such bones usually

form when the lambdoidal

. . is under stress, as might result

suture.

from powerful nuchal action drawing the occiput downward”. Parietal II preserves 29 mm of the squamous suture immediately posterior to the pterion (which is missing), showing some ridges for the articulation

temporal

squama/parietal

bone.

The five fragments composing Parietal II converge towards the parietal protuberance, being broken near it. The external surface is in good condition. The fragment AT-21 1 exhibits increases

a brief segment from the sutures

lambdoidal vicinity.

borders)

The curvatures the middle border

towards

of Parietal

of the bone.

of AT-61.

very shallow,

(28 mm) (varying the

of the temporal between 6 and middle

eminence,

II are similar

A small

reaching

abrasion

roughly

circular

some bone condensation

circumscribing

the lesion. The origin

with a subsequent

inflammation.

There

weakly

marked

Alongside diploic

and capillary the preserved

networks sagittal

holes are discernible

The

impressions

of 10 mm

near

located

the

in

the coronal and

looks smooth

of this pathology

but

in its

1.2 mm of diameter) surface

bone and

with

is traumatic

(P. J. Perez, parietal,

personal

frontal

and

as well as some main fissures and gyri (Figure 5). is even and no cerebral impressions are present.

are also present

suture

abraded

are no signs of infection

cerebral

temporal lobes can be distinctly discerned On the contrary P. I endocranial surface Anastomosis

table.

of the sagittal

I, with theparielum

(with some

only the outer

communication). Parietal II shows

bone

a value

can be observed

affecting minor

The thickness in the coronal,

to those of Parietal

pathological

It is well delimited,

lines. 8 mm

in P. II but not so developed

the left side of the sagittal

along the sulcus. Near the obelion,

as in P. I.

sulcus can be seen. Several

8 mm from the sagittal

suture,

there is a rounded and shallow depression corresponding to a Pacchioni’s fossa, with many small diploic foramina. In the wormian bone AT-76 there is a blunt ridge or buttressing at the midline. Parietal III (P. III). This specimen both

found

in 1984) constituting

(Figure 3). It preserves 27 mm) and a segment

consists

of two parietal

the posterior

the parietomastoid suture (53 mm) of the lambdoidal

inferior

fragments

quadrant

(AT-63

of a right

and AT-65, parietal

bone

eroded and lacking indentation (length = border. The superior halfof the preserved

lambdoidal suture shows complete internal obliteration and a very advanced degree offusion on the outer table. Ridges are present at the parietal notch, and the striaeparietalis appear, as in Parietal I and Parietal II, weakly developed. The thickness at the asterion is of 9.5 mm in the suture and 11.5 mm in the ending of the superior temporal line at the mastoid border (near the asterion). The temporal lines are very tenuous and there is no angular torus. Parietal

III articulates

with the occipital

fragment

AT-86 (the set is called Cranium 1) The meningeal vascular system is represented

AT-122

and with the temporal

in P. III by a lambdatic

main collateral vessels (Figure 6). The most inferior of them is narrow ending in a Pacchioni’s fossa close to asterion. There is also in P. I a similar of the lambdatic ramus which pours in a Pacchioni’s fossa.

branch

fragment with four

and sharp edged collateral branch

CRANIAL

REMAINS

AND

LONG

BONES

FROM

203

ATAPUERCAIIBEAS

3. Occipital bone Occipital I(0.

I). This specimen consists offive left side occipital fragments:

AT-106

(found

in 1985), AT-2 15 ( 1988)) AT- 105 ( 1985) and AT- 132 ( 1986). Occipital I preserves 64 mm of left lambdoidal suture and 28 mm of left occipitomastoid border (Figure 7). Towards the midline 0. I extends

as much as 68 mm from the left asterion.

semispinalis capitis (or m. complexus) is well preserved

The impression

of the m.

and there is some buttressing

of the

superior nuchal line above it. The thickness ofthe nuchal ridge increases towards the midline, reaching a maximum

of 11 mm at the medial break. The ridge does not extend laterally to the

m. semispinalis capitis. The inferior nuchal line is well marked due to the marked excavation the m. obliquus superior impression.

It is interesting

to note that the maximum

of

thickness of the

specimen is not located at the occipital torus but in the transverse sulcus (at its medial break = 14.5 mm). The asterionic thickness is 9 mm. The transverse sinus appears medially as a sulcus but this sulcus vanishes towards the asterion, persisting as a slight butressing that separates the cerebral and cerebellar fossae. This atypical pattern is found in all the occipital asterionic

regions of the Atapuerca’s

hypodigm:

0. II, 0. III,

AT-122,

AT-123a/b

and

AT-39. Occipital II (0. II). F ive occipital fragments are joined in 0. II: AT-45 (found in 1984), AT-56 ( 1984) and AT-20 1a/AT-20 1b/AT-20 1c ( 1988) (F’g I ure 7). The complete right lambboidal border is preserved (lambda-asterion chord = 85 mm) as well as segments of the left lambdoidal suture (extending 25 mm from the lambda) and right occipitomastoid border (extending

25 mm from asterion).

Only the most lateral end of the right m. semispinalis capitis impression is preserved in 0. II, and there is no buttressing of the superior nuchal line above it. So, as in 0. I the nuchal ridge do not reach the lambdoidal suture at the asterion. The thickest points of the specimen lay on the sagittal sinus (11 mm) and transverse sinus (11 mm). The lambda thickness is 9.5 mm and the asterion thickness 7.7 mm. The internal surface of the lambdatic

fragment

instead of the superior sagittal sulcus commonly Occipital

III (0. III). The specimen

1987), AT-206

(found in 1988), AT-40

AT-201b

consists of three fragments: (1984) and AT-216

1988). This is the best preserved occiput in the Atapuerca left asterion (Figure 7). The left lambdoidal lambda

and asterion

(lambda-asterion

bears a blunt midsagittal

crest

found. AT-177

(unearthed

in

(found in 1987 but identified in

hypodigm,

preserving lambda and

border is severely eroded except in the vicinity of

chord =88 mm).

The

right lambdoidal

suture is

preserved for 34 mm ‘from lambda, showing advanced obliteration, and the left occipitomastoid border extends for 30 mm from asterion. A great part of the occipital plane is present in 0. III, especially the left half. The nuchal ridge is preserved in two disjoint segments. One of them (very altered)

extends 54 mm from left asterion

medially.

The other one extends

20 mm from the midline to the right; at this level a marked angulation between the occipital and nuchal planes can be observed. The lambda-inion chord and arch can be taken at inion or just lateral

to it. A chord of 63.5 mm and an arch of 69.5 mm are estimated

following

Hublin’s (1978a) recommendations for the placement of inion (i.e., on the inferior border of the occipital torus at the midline). Comparing with Hublin’s (1984) data, 0. III, Swanscombe and La Chaise (both juvenile and adult occiputs from Abri Suard) fall within the range of 12 upper Pleistocene Neandertals from western Europe. In this specimen the morphology of the occipital plane can be determined: the curvature is regular without occipital bunning or lambdoidal flattening and the suprainiac fossa absent.

204

J. L. ARSUAGA

ET AL.

CRANIAL

REMAINS

205

AND LONG BONES FROM ATAPUERCA/IBEAS

Figure 6. Endocast ofParieta1 III+AT-122 (occipital fragment). A = Asterion. lb = Lambdatic branch of the meningeal vascular system. LSS = Lobulus semilunaris superior (cerebellum). OG = Occipital gyri. PF = Pacchioni’s fossa. PI = Parietal incisure. SS = S&us lateralis.

Twenty-eight

millimetres

below

lambda

there

is a depressed

traumatic origin (P. J. Ptrez, personal communication). The thickness ofthis specimen is noteworthy reaching a maximum

small

area

of probably

of 22 mm in the nuchal

ridge near the midline. The thickness at lambda is 9 mm and 11.5 at asterion. 0. III preserves almost completely the path of the sinus sugittulis superior from lambda to the cruciate eminence (eminentia cruciutu). The saggittal sulcus is well developed inferiorly but changes to a buttress-like appearance towards lambda. The sagittal sulcus continues directly into the right transverse sulcus in the preserved portion.

AT-122. (found in 1986). The specimen includes the right asterion, 44 mm of right lambdoidal suture and 38 mm of right occipitomastoid suture, extending 40 mm medially (from asterion). AT-122 articulates with Parietal III (Cranium 1) along its lambdoidal border. Only the most lateral part ofthe m. complexus impression is preserved. The superior nuchal line is not buttressed above it, but seems to continue as a slender ridge towards the asterion. The maximum thickness of the fragment (12 mm) lays on this ridge.

AT-39 (recovered

in 1984). This specimen is roughly rectangular with 24 mm of the right lambdoidal suture and 35 mm of the right occipitomastoid border is preserved and intersects at the asterion. The fragment extends medially some 32 mm from asterion. The m. complexus insertion is completely

missing and there is no sign ofthe superior nuchal line, so it disappears

206

J. L. ARSUAGA

Figure 7. Occipital bones. (1) Occipital

ET AL.

III; (2) Occipital

II; (3) Occipital

I.

CRANIAL

REMAINS

AND LONG BONES FROM ATAPUERCA/IBEAS

long before reaching the asterion. The maximum thickness of the specimen the transverse sinus. The thickness at the asterion is only 6.5 mm.

207

(11.5 mm) lays in

AT-66 (found in 1984). Roughly rectangular, this fragment preserves 20 mm of lambdoidal border and 24 mm of the suture connecting AT-66 with the wormian bone AT-76 (Cranium 2). Another much smaller extra-sutural bone (missing) was situated in the right lambdoidal suture between AT-66 and AT-76. The maximum thickness of AT-76 is 11 mm. AT-123a/AT-123b (foun d in 1986). This specimen preserves the right asterion, 17 mm of lambdoidal suture and 35 mm of occipitomastoid border. It extends medially 40 mm from asterion. The external surface is severely damaged but seems like Occipital III. The maximum thickness of the fossil is IO.4 mm ( 10 mm at asterion). 4. Temporal bone (Figure 8) AT-84 and AT-86. These two temporal remains represent almost analogous regions from different sides and individuals (Martinez & Arsuaga, 1985). AT-84 (found in 1984) is part of a left temporal bone. The mastoid region is largely complete with some minor loss of bone at the tip of the mastoid process. Only the squamous portion which extends behind the supramastoid crest is preserved. Medially to the internal acoustic meatus the petrous bone is absent. There is also a small portion of the tympanic plate. AT-84 preserves the totality of the parietomastoid suture (30 mm) and the occipitomastoid suture (58 mm). The bone thickness at asterion is 8.5 mm. AT-86 (found in 1984) belongs to a right temporal bone. The mastoid region is only represented by the mastoid process. The external acoustic meatus and the posterior root of the zygomatic process are present. The squamous portion is missing except for a small region including the parietal notch which articulates with Parietal II (Cranium 1). Most of the tympanic plate is preserved as well as the portion of petrous bone laterally to the internal acoustic meatus. In AT-84 and AT-86 the mastoid processes are oriented to inferior. They are wide at the base and in AT-84 the process tapers toward its tip. Both processes look very projecting (Table 4) and well isolated from the surrounding bone. AT-84 exhibits a prominent mastoid crest running on the posterolateral surface of the mastoid process. A deep supramastoid sulcus is discernible between the mastoid crest and the preserved portion of a strong supramastoid crest. AT-86 displays weaker mastoid and supramastoid crests as well as a more shallow sulcus between them. The supramastoid crest does not overhang the external acoustic meatus neither in AT-84 nor in AT-86. The anterior mastoid tubercle (sensu Hublin, 19786) is absent in both temporal bones. The mastoid notch is preserved in AT-84 coursing toward the stylomastoid foramen. AT-84 shows a developed juxtamastoid eminence (sensu Rouviere, in Hublin, 19786) (Table 4), separated from the occipitomastoid suture by the s&us arteria occipitalis.

On the endocranial side ofATand AT-86 the sigmoid sulcus can be recognized but it is complete only in AT-84, entering in the temporal bone below asterion. The petrous bone is well preserved in AT-84 and AT-86. On thefacies cerebralis several anatomical structures can be studied, mainly the hiatus canalis nervu.sfacialis, the eminentia arcuata (very distinct in AT-84) and part of the tegmen tympani. In AT-86 and particularly in AT-84, the anterior surface of the pyramis is uneven because of the presence of the gyrus occipitotemporalis lateralis. On thefacies cerebellaris both pieces preserve theporus acusticus internus (complete in AT-84 and only the lateral margin in AT-86)) thefossa subarcuata (petromastoid canal sensu Gannon et al., 1988) and the apertura externa aqueductus vestibuli (sen.w Weidenreich,

208 Table 4

J. L. ARSUAGA

ET AL.

Temporal bone meamuremaats (mm) 1M

Sample

Sepdlveda (M) ’ Average S.D. Jv Range Seplilveda (F) ’ Average S.D. x Range Shanidar?

4.7 1.9 24 0.5-10.5

44.7 4.3 26 35.5-53.5

24.4 2.4 34 2&32

11.2 2.5 34 6-15.5

3.3 1.6 34 0.5-7

38.5 4.4 33 30.546.5

8.3 10.6

11.3 6.6

__

(&9) /Ibeas’

4M

13.9 3.1 27 7.5-18.5

1

38-2 1 Atapuerca AT-84 AT-86

3M

26.6 2.3 26 22-31.5

2 West European Neandertals La Ferrassie I3 27 La Chapelle-auxSaints3 28 Gibraltar4 23.4 La Quina H!+ 24 La Quina H 1O3 25 La Quina H273 26 SPY l3 27 SPY 23 26? Krapina4 C 22.4 39- 1 22 38-7/38-l 1 27.4 38-2/38-14 28.3 38-12 23 38-13 39-14

2M

(21) 29.5 23

6 10 3.2 7 6.5 6.5 7 7 4 3 7.2 9.7 4.6 5.8 IO.4 4.4 14 11

8

1 M: Length of the mastoid process base (method of Zoja, in Vallois, 1969). 2M: Projection of the mastoid process (method ofZoja, in Vallois, 1969). 3M: Height ofthe eminentiajuxtamasfoi (mastooccipital crest for Shanidar 1 & 2) measured from the deepest point in the digastric sulcus (Trinkaus, 1983). 4M: Distance from the incisurapnridalis to the tip of the mastoid process. ‘Measurements taken by the Authors. ?From Trinkaus (1983). 3From Vallois (1969). *From Smith (1980). ( ) Measurements in parentheses are minimum measurements. (M) Males. (F) Females.

1943)) which opens into a deep impressio cerebellaris. In AT-84 and AT-86 the margo superior of the pyramis bears the sulcus superior petrosal sinus. In AT-84 the posterior surface ficies cerebellaris) is overhung by a sharp crista pyramidis. In AT-86 this crest is rounded but the posterior surface is concave. According to Weidenreich (1943) the “Sinanthropus” temporal bones shows a different pattern, with a rounded cristapyramidis and without a concavefacies cerebellaris. In the fucies inferior of both Atapuerca temporal bones parts of the lateral wall of

CRANIAL

REMAINS

AND LONG BONES FROM ATAPUERCA/IBEAS

209

the carotid foramen are preserved as well as the stylomastoid foramen, the base of the styloid process, the fossula petrossa and most of the jugular incisure. Only a small portion of the tympanic plate (including a well developed vagina processus styloidei) is preserved in AT-84 whereas in AT-86 the totality of the tympanic plate laterally to the sulcus anuli Qmpanici is present. In AT-86 a thick and blunt tympanic crest runs from the styloid process to the middle of the inferior margin of the external auditory meatus. Therefore, the tympanic plate seems divided into two sides by the crest. There is also another small crest extending from the styloid process to reach the margin of the external auditory meatus at its most posterior point. The external auditory meatus is circular. AT-124 (found in 1986). This specimen comprises a left glenoid fossa and surrounding regions including the root of the zygomatic process, most of the squamous wall of the external auditory meatus and small portions of the tympanic and petrous bones. On the lateral face the root ofthe zygomatic process strongly projects laterally and forms a wide ( 10 mm) sulcus processus zygomatici (sensu Weidenreich, 1943). The acoustic porus lays below the zygomatic process and also below the roof of the mandibular fossa. The postglenoid process is strikingly developed. A part ofthe infratemporal area with the sphenoid margin can be identified in basal view. The anterior wall of the mandibular fossa appears deeply concave from side to side and there is a smooth transition between the articular joint and the preglenoid planum. So, the articular tubercle is not raised. Signs ofa temporomandibular arthrosis, which did not substantially alter the original morphology, can be seen in AT- 124 (Perez & Martinez, 1989). A strongly developed postglenoid process forms the posterior wall of the mandibular fossa, extending medially to the vicinity of the well developed entoglenoid process. AT-220 (found in 1986 but identified in 1988). This fossil is a small portion ofa right petrous bone. Laterally to the internal acoustic meatus thefacies cerebralis is preserved with the hiatus canalis nervus facialis and a portion of the eminentia arcuata. In the facies cerebellaris the fossa subarcuata and the apertura externa aqueductus vestibuli (sensu Weidenreich, 1943) are present. The margo superior is unsharpened. Rests of the carotid foramen, jugular incisure andfossulapetrosa are preserved in the facies inferior. The cochlea is exposed by fracturing. AT-125 (found in 1986). This specimen corresponds to the mastoid angle of a right temporal bone, preserving the totality of the parietomastoid border (27 mm) and part of the occipitomastoid border (30 mm). The thickness at asterion is 6-5 mm. AT-125 shows, as AT-220, an extensive pneumatization. On the endocranial surface the sigmoid sulcus enters in the temporal bone below asterion. 5. Humeri: inventory, state of preservation and description

AT-25 (found in 1984). This specimen corresponds to the proximal half of a right humerus (maximum length = 147 mm). The humeral head is damaged, with substantial bone loss posteriorly (about one-fourth ofits mass). The lesser tubercle and greater tubercle also show some superficial erosion. The diaphyseal surface is well preserved and the muscular insertions appear poorly developed. The intertubercular sulcus looks straight and very broad. The anatomical margins are blunt. AT-25 values for several humeral measurements are shown in Tables 5 and 6. It can be observed that the AT-25 figures fall within the ranges of the Neandertal sample. Neverthe-

210

ET AL.

J. L. ARSUAGA

Measurements

Table 5

of the Atapucra

Length of fragment

humeri

(mm)

AT-25 170

AT-93 146

AT-2 17 138

S.N. level Perimeter Maximum diameter Minimum diameter P.M. level Perimeter Maximum diameter Minimum diameter D.T. level Perimeter (A) Maximum diameter Minimum diameter Width oftuberosity (B) Relative width B/A (%) M.S. level’ Perimeter Maximum diameter Minium diameter Shaft index Anterior cortex Posterior cortex Medial cortex Lateral cortex Cortical area (mm) Articular head Vertical diameter Transverse diameter Index Neck angle

85 30 19.6

87 30.7 25.3

73 24.5 21

81.5 27.7 24.7

72 24.3 19.6 15 20.8

82 28.6 26 20 24.4

70 24.5 18.8 76.7 6.5 7 6 6 340

27.7 (18.8) 67.8 7.5 6 5.5

74 25 21 84 8 5.5 5.5 5 330

45 >45 > 100 135

S.N. = Measurements taken at surgical neck level. P.M. =Measurements taken at midpoint of m. Pectoralis Rlnjor insertion area. D.T. = Measurements taken at midpoint of Deltoid tuberosity (near 5/12 of maximum length level). M.S. = Measurements taken at midshaft. ’ = The Atapuerca specimens are broken at a level short to midshaft.

less, there is a broad overlap in humeral diameters between Neandertals, and a Spanish Mediaeval sample from Septilveda. Remnants

of epiphyseal

closure are visible in AT-25.

Upper Palaeolithics

The age at death of the specimen

would be around 25 years according to modern human patterns (Schranz, 1959; Olivier, 1960, Krogman & Iscan, 1986; Bass, 1987). The structure of the cancellous tissue of the proximal epiphysis and the extension of the medullary cavity corresponds to Phase I as defined by Acsadi & Nemeskiri ( 1970) (Figure 9). AT-93

(discovered

in 1976). This right humeral

fragment

extends from the proximal

epi-

physeal line level to the deltoid tuberosity distal end (maximum length= 146 mm). The surface is in generally good condition. The intertubercular sulcus is preserved almost completely, appearing (as AT-25) straight and broad. The muscular attachments are well developed in AT-93,

much more than in AT-25.

AT-93

measurements

are greater than in AT-25

CRANIAL

Table6

Mess -ads

211

REMAINS AND LONG BONES FROM ATAPUERCA/IBEAS

of the humeri (mm) Humeral shaft

Midshaft perimeter

Humeral head

Deltoid tub. perimeter

Diameter max. midshaft

Diameter min. midshaft

Midshaft index

72 82

24.5 27.7 25

18.8 18.8 21

76.7 67.8 84

13 66.7 5.2 58-74

16 23.1 2.3 20-26.8

16 16.4 1.7 14-19

16 71 3.6 65.2-74.5

12 2168 19-25.5

12 1746 1620

12 80.77 72.5-94.7

43 21.8 17-27.6

43 17.4 14-20.6

43 79.3 70.2-93

AT-25 70 AT-93 AT-2 17 74 Neandertal Parameters’ N 15 Mean 65.8 SD. 6 Range 56-77.5 European Upper Palaeolithicr (only right side) N Mean Range Seplilveda Mcdiaevals3 N Mean Range Adult Whites’ Mean 0 (&= 200) Mean 0 (N= 200) Difference

Vertical diameter

Transverse diameter

Width index

245

r 100

39 41.9 34.1-48

35 39.7 33.245

34 94.7 87-100

48.76 42.67 6.09

44.60 36.98 5.68

45

Deltoid tub. = Deltoid tuberosity. max. = maximum. min. = minimum. I = Measurements of La Ferrassie and La Chapelle by the authors. Other Neandertals: Endo & Kimura (1970); Thoma (1975); Lovejoy & Trinkaus (1980), Trinkaus (1983). X represents the number of humeri available for measurement (one or two per individual). 2= Measurements from Thoma (1975). ’ = N represents the number of measured humeri because it is not possible to determine individuals in the Sepulveda bone accumulation. ‘= Measurements from Dwight (1904-1905) in Krogman & Iscan ( 1986).

(Table

5). The perimeters

Neandertal The

average (Table

trabecular

system

at deltoid tuberosity

and midshaft levels are noticeably

above our

6). and

extension

of medullary

cavity

match

with

Acsidi

&

Nemeskeri’s Phase I (Figure 9). The proximal break of AT-93 is largely coincident with the proximal epiphyseal line. Signs of recent epiphyseal closure can also be recognized. So, AT-93

and AT-25

show similar degrees of bone maturity.

AT-217 (found in 1988). This right humeral

specimen

extends from the midshaft

to just

above the medial epicondyle (maximum length = 138 mm). The distal break runs diagonally and only the medial pillar surrounding the olecranon fossa is preserved (Figure 9). The external surface is well preserved, and there is no nutrient foramen. The three margins of the bone are dull and the three surfaces appear fairly convex. The proximal cross section shape is oval. 6. Humeri: morphology in detail and comparisons Deltoid tuberosity. In AT-25 the deltoid tuberosity, anterior

margin

and the other one laterally

weakly

marked,

has one crest on the

placed. The sulcus between

these crests looks

212

1. L.

ARSUAGA

ET AL.

Figure 8. Temporal bones. (I ! AT-86: Posterior view; (2) AT-W Posterior view; (3) AT-86: Latrral (4) AT-84: Basalview: (5) Al’-124: Latcral view: (6) AT-124: Basal view.

view;

CRANIAL

REMAINS AND LONG BONES FROM ATAPUERCAIIBEAS

213

214

ET AL.

J. L. ARSUAGA

shallow. On the other hand, the tuberosity is strongly marked in AT-93 between the anterior

and lateral ones. Proximally

does not reach the external margin of the humerus. This morphology not reaching Mediaeval

the external

margin)

sample from Seprilveda

and has a third crest

the deltoid tuberosity

was found in two humeri

in both specimens

(i.e. deltoid tuberosity

out of 43 in the Spanish

that we have used for comparison.

According

( 197 1)) the deltoid tuberosity is narrower in “Sinanthropus” and “classic” Neandertals West Asian Neandertals

and recent humans. In the Atapuerca

width is greater than in “classic” Asian Neandertals,

Qafieh

Neandertals

to Endo than in

specimens the relative deltoid

(Table

6), and it is similar to those of West

specimens (Vandermeersch,

1981) and recent humans. Another

feature considered by Endo ( 197 1) as typical of Neandertals (both “classic” and West Asian) is a deltoid tuberosity with two crests, while the recent Lebanese sample studied by Endo displayed three crests as a rule. However, in the Spanish Mediaeval sample from Septilveda we found 14 specimens with two crests out of 41 humeri. Therefore, as other authors have pointed

out (Thoma,

discriminating

1975; Vandermeersch,

character.

Furthermore,

1981) the presence

of a third crest is not a

AT-25 exhibits two crests and conversely AT-93 has

three crests (Figure 9). Surface medial to the deltoid tuberosity. According this surface is distinctly flat or concave.

to Thoma

convex in Neandertals,

In AT-25

( 1975) and Endo & Kimura

whereas in modern populations

the surface medial to the deltoid tuberosity

looks broad and concave.

This character

state (flat or concave

is Aat and in AT-93

surface)

is found in every

humerus from SepGlveda, and in the two humeri from Qafzeh (Vandermeersch, Humeral head. In modern populations

the vertical diameter

( 1970)

it would be

of the humeral

198 1) .

head is greater

than the transversal diameter and so the head looks transversally compressed (Krogman & Iscan, 1986). In AT-25 the humeral head index can be estimated in 100 or more and the humeral

head appears

the humeral

head index is 106 (Basabe,

Dusseldorf

rounded

or even transversally 1966)) reflecting

oval (Table a rounded

5). In Lezetxiki

humeral

head, as in

(Boule, 1911-13).

Lesser tube&e. The morphology the Seplilveda antero-medially

Mediaeval

of the AT-25

lesser tubercle is clearly different in shape from

sample. The lesser tubercle is massive, very broad at the base and

protruded.

In the modern humeri from SepGlveda the lesser tubercle looks

slender. Also in AT-25 the area between the lesser tubercle and the anatomical neck (related to the glenohumeral superior ligament) is very narrow with respect to modern humeri. In AT-93 the preserved portion of the lesser tubercle looks like AT-25. Trinkaus ( 1983) reports a large and projecting lesser tubercle in Shanidar 1. The massiveness of this structure is related to a powerful m. subscapularis. The area of insertion of the gleno-humeral inferior ligament constitutes

a very depressed triangular

fossa in AT-25

(so the ligament would be very devel-

oped). In the Septilveda sample this area never forms a fossa. The morphology of the* preserved portion of the glenohumeral inferior ligament attachment in AT-93 is similar to that ofAT-25. Cortical thickness. AT-25

and AT-217

are broken

near midshaft.

The

Atapuerca

humeri

display at this level much thicker cortical area than any of the Seplilveda specimens (Figure 10). The cortical area (Table 5) represents 85% of the total area in AT-25 and 73.3% in AT-217. The Seplilveda average is only 56.7% (Figure 11). All the AT-25 and AT-21 7

CRANIAL

REMAINS

AND

LONG

BONES

FROM

215

ATAPUERCA/IBEAS

TA, CA, ACPC. -

AT-25

----

AT- 217

MC* LC,

Figure 10. Cortical thickness of the Atapuerca/Ibeas humeri AT-25 and AT-217 at midshaft level. The cortical thickness of the Atapuerca humeri has been compared with that of 15 modern humeri from the Seplilveda sample which were cut at midshaft. Photographs of the cross sections were taken and the anteroposterior axis drawn as the line passing through the centre of the medullary canal and the anterior margin of the humerus. The mediolateral axis was taken as a perpendicular to the anteroposterior axis through the centre of the medullary space. TA=Total area (mm); CA= Cortical area (mm); AC= Anterior cortex (mm); PC = Posterior cortex; MC = Medial cortex; LC = Lateral cortex. The values ofthe Atapuerca/Ibeas humeri have beenstandardized with respect to the modernsampleofSepulveda (.N= 15). SD =standard deviation.

%

10

70

AT-217

ML+--_

80

90

AT-25

* AT-217

CA

AT-25

a

ITA --hT AT-217

AT-25

Figure 11. Three indexes of cortical thickness for the Atapuerca/Ibeas humeri at midshaft level. AP: Anteroposterior cortical index = (anterior+ posterior cortex x lOO)/anteroposterior diameter. ML: Mediolateral cortical index = (medial + lateral cortex x lOO)/mediolateral diameter). CA/TA = cortical area x lOO/total area. The Atapuerca/Ibeas values are compared to the modem sample of Sepulveda (N= 15). Range and mean f 1 SD of the Septilveda sample are indicated.

216

ET AL.

J. L. ARSUAGA

measures and indexes of cortical thickness are more than one standard deviation above the Sepulveda mean (Figures 10 and 11). Also, according to Ben-Itzhak et al. (1988) the cortical thickness

(both anteroposterior

significantly

and mediolateral)

of Neandertal

right and left humeri are

higher than in modern Homo sapiens.

Ridgefor the lateral head of m. triceps. According this ridge is absent in Neandertals. sample) it appears frequently.

Conversely,

to Endo & Kimura

(1970) and Thoma

in modern humans (including

This crest is present in AT-93

( 1975)

the Sepulveda

but not in AT-25.

7. Ulna AT-218 (found in 1988). Proximal fragment of right ulna lacking proximal epiphysis except the radial facet (maximum length = 44 mm). The proximal break exposes a coarse travecular system and the distal break exhibits a circular cross section with thick cortical bone. The crista m. supinatoris is noticeably broad and blunt in contrast with modern people. The AT-218 radial facet appears (judging from the preserved portion) small and rounded, like in Ferrassie 1, Shanidar

4 and the Krapina

small or almost absent in Amud (Thoma,

1975). In AT-218

sample. The hollow for the play of the radial tuberosity 1, Dusseldorf

(Endo & Kimura,

is

1970), Spy 1 and Spy 2

the hollow is also absent.

8. Tibiae AT85 (found in 1984). This fragment of right tibia extends from the proximal end of the tibia1 tuberosity to the midshaft (maximum length = 167 mm). The metrical variables of the fossil are presented in Tables

7 and 8. The external surface is in good condition.

The shaft shows a small curvature in the transverse direction. The nutrient foramen is completely preserved and there is another (much smaller) foramen above it (Figure 12). The anterior margin is round. In modern tibiae the anterior

margin passes laterally to the tibia1

tuberosity but in AT-85 this margin merges into the tibia1 tuberosity. The internal margin is round and blunt. It is located in the middle of the medial side of the tibia, like in Amud and La Chapelle-aux-Saints,

and not in the posterior side as commonly

The external margin is weakly marked. It is interesting

occurs in modern tibiae.

to note that the well developed soleal

line does not reach the medial margin. The posterior margin above midshaft.

(vertical ridge) is upstanding.

This feature is absent in La Ferrassie

It runs from the soleal line and ends 1 and 2 and La Chapelle-aux-Saints

and it is not common in the Sepulveda sample (14 tibiae out of 79). Endo & Kimura reported a projected verical ridge in Amud 1.

( 1970)

The cross section shape at the midshaft is amygdaloid. This shape has been described in several Neandertal specimens (Lovejoy & Trinkaus, 1980; Trinkaus, 1983). Trinkaus (1983) contends that the amygdaloid exceptional robusticity.

AT-19 (found in 1976).

cross sectional shape in Neandertals

This specimen

is a fragment

is associated with

of right tibia extending

from the

midshaft level to the least circumference level (maximum length = 107 mm) (Figure 12). The metrical variables of AT-19 are presented in Tables 7 and 8. The external surface is well preserved. The three anatomical margins are blunt. The internal margin is also located more anteriorly than is common in modern tibiae, as in AT-85. The three surfaces are convex. The shape of the cross section at the midshaft is amygdaloid as in AT-85.

CRANIAL

Table 7

REMAINS

AND

LONG

Meamuemeats of the Atapuera

Length of fragment

Foramen nutricial level Perimeter A-P diameter M-L diameter Cnemic index Midshaft level’ Perimeter A-P diameter M-L diameter Midshaft index Anterior cortex Posterior cortex Medial cortex Lateral cortex Cortical area (mm) 2/3 Total length level Perimeter A-P diameter M-L diameter Anterior cortex Posterior cortex Medial cortex Lateral cortex Cortical area (mm)

BONES

FROM

ATAPIJERCAIIBEAS

217

tibiae (mm) AT-85 167

AT-19 107

TB-I 125

93 36.3 24 66.1 80 29.3 22.5 76.7 9.5 7.5 4.5 5 340

76 27.6 20 72.4 9.5 6.5 4.5 4 320 84 29.5 24.5 7.5 8 5.5 6 400

A-P = Anteroposterior. M-L = Mediolateral. ’ = AT-85 & AT-19 are broken at a level short to midshaft.

Tibia I. The specimen corresponds to the distal third of a left tibia (Figure 12) and consists of two fragments that completely match: AT-91 (discovered in 1984) and AT- 119 (found in 1985). The external surface is in good state of preservation except above the distal articular surface where the cortical bone is mostly missing. The medial malleolus is also lacking and the two projections of the fibular articular surface are eroded. The three margins of the diaphysis are round and blunt and the surfaces convex. The cross section at the proximal break is amygdaloid. In our modern sample (Sepulveda) the cross section looks subtriangular. The talar trochlear surface is flat in respect to the Seplilveda tibiae. Both the anterior and posterior margins do not project distally. The ridge which runs sagittaly across the middle of the articular surface is low and not clear (Figure 12). These traits are also reported in Amud 1 (Endo & Kimura, 1970) and Spy 2 (Thoma, 1975) and we have observed them in La Ferrassie 1 and 2. The fibular articular surface is shallow as in Amud 1 (Endo & Kimura, 1970) and La Ferrassie 1 and 2. Tibia I does not exhibit a squatting facet. This trait is present in some Neandertals (Trinkaus, 1975, 1983) as well as in modern populations (Carretero et al., 1987; Perez et al., 1988). The phylogenetic relevance of this facet is null. Cortical thickness. AT-85 and AT-19 are broken more or less at midshaft level and TB-I at 2/3 of total length level. At midshaft level AT-85 and AT- 19 differ from our modern sample only

level

7

21.64 175-25

105 23 17.2-31

7 37.90 32-45

105 31.8 23-40.6

105 71.6 58.4-88

12 72.3 6.9 62-75

Cnemic index

6 84 74100

11 88.3 7.5 75-90

80 76

Perimeter

6 20. I 7 17-24

100 21.1 17-30

100 27.9 19.5-34.3

13 22.8 1.6 20-25.8

22.5 20

M-L diameter

6 32.25 27-39

13 32.5 3.6 27738.4

29.3 27.6

A-P diameter

Midshaft level

100 76 52-94

6 63.22 50-7 1.4

13 70.4 4.7 626-77

76.7 72.4

Midshaft index

70.5 53-83

99

7 76.43 62-89

67 84

Least perimeter

5 41.90 33-60

8 37.5 3.0 31.541

(40)

A-P diameter

92 46 37-54

4i.40 44-55

8 50.6 4.7 44-54

(53-55)

M-L diameter

Distal epiphysis

’ = Measurements

A-P = Anteroposterior. M-L = Mediolateral. ’ =Sources as in Table 5. .,V represents the number of tibiae available for measurement (one or two per individual). from Thoma ( 1975). Only one side per individual is represented. 3= Mrepresents the number of measured tibiae because it is not possible to determine individuals in the Seplilveda bone accumulation.

12 26.8 1.7 2429.6

12 37.4 4.1 28842.6

7 105.4 3.3 101-l 12

24

M-L diameter

nutricial

36.3

A-P diameter

Foramen

of tibiae (mm)

93

Perimeter

MePsuremcmts

AT-85 AT-19 TB- 1 Neandertal Parameters’ JV Mean SD. Range European Upper Palaeolithic’ “V Mean Range Seplilveda Mediaevals’ “Y Mean Range

Table 8

CRANIAL

REMAINS

AND

LONG

BONES

FROM

ATAPUERCA/IBEAS

219

220

J. L. ARSUAGA

-1

- -l!

ET AL.

II

1

112 TA CA

>

>

NM

MC LC -

-

-

-

--

213 TA

*.._ a. CA

. ..*

..

. . ..I

AC

. . .. ‘....

... . .. . . . . .

PC

,.*.-*

. ...a.

*

-

AT-19

----

AT-05

.*. . . * -.. *.

MC LC -

-

-

A-

Figure 13. Cortical thickness of the Atapuerca/Ibeas tibiae at midshaft level (l/2) and 2/3 of total length level (2/3). Methodology and abbreviations as in Figure 10. The values ofthe Atapuerca/Ibeas tibiae have been standardized with respect to the modern sample of Seplilveda (N= 15).

in a thicker posterior

wall (Figures 13 and 14). In TB-I, at 2/3 level, the posterior cortex is again much thicker than in the Sepulveda sample (more than six standard deviations above the Seplilveda mean!). The medial and lateral walls ofTB-I are also very thick and therefore the cortical area is great both in absolute and relative terms (Figures 13 and 14). Several

studies suggest that the cortical humans (Endo & Kimura,

bone in Neandertal

1970; Lovejoy & Trinkaus,

tibiae is thicker 1980; Trinkaus,

than that of modern 1983). Thick-walled

long bones have also been reported for African and Asian Homo erectus (Kennedy,

1983, 1985;

Day, 1971).

Discussion

and conclusions

In this section a phylogenetic analysis ofcharacters is attempted. Only some of the previously described traits are useful for this purpose. To establish the polarity (i.e., direction of evolutionary change) of characters, early and middle Pleistocene African and Asian fossils have been used as outgroups.

CRANIAL

%10

REMAINS

AND

LONG

BONES

FROM

30

20

1,

v2

70 4p

y

5p

221

ATAPUERCA/IBEAS

BO

90

+

AT-19

ML+ AT-95

CA JTA

4

m-19

213

AP -16-I ML--J--* TB-I

CAITA .--A TB-I Figure 14. Three indexes ofcortical thickness for the Atapuerca/Ibeas tibiae at midshaft level (l/2) and 2/3 of total length level (2/3). Abbreviations as in Figure 11. The Atapuerca/Ibeas values are compared to the modern sample of Seplilveda (X- 15). Range and mean + 1 SD of the Sep6lveda sample are indicated.

Concerning the morphology of the orbital and lateral segments of the supraorbital torus, we recognize three character states in Homo erectusltlomo sapiens. The supraorbital torus (including the lateral segments) looks straight in anterior and superior views in the Zhoukoudian sample, in the Javanese specimens from Sangiran, Trinil, Sambungmachan, Perning and Ngandong as well as in the African crania KNM-ER 3733, KNM-ER 3883 and OH 9 (observations on casts). We believe that this morphology represents the plesiomorphic condition. On the other hand there is no straight junction of the torus and frontal squama in the middle Pleistocene African and European “late Homo erectus” or “early Homo sapiens”, as well as in Neandertals. In these fossils the supraorbital trigone (lateral segment of the supraorbital torus) is deflected downward. This is the character state present in AT-121. Finally, in anatomically modern humans the torus is very much reduced in thickness and projection. So, with respect to this character the European middle Pleistocene fossils and the Upper Pleistocene Neandertals share the same state. Nevertheless, the morphology of the glabellar segment of the torus is clearly autapomorphic in Neandertals (from either the Riss-Wiirm or the Wiirm). In this group the superciliary arches and the glabella are so completely fused that the supraorbital torus appears continuous in the glabellar region. AT-200 exhibits this Neandertal apomorphy. In other human fossils assigned to Homo erectus, archaic Homo sapiens or anatomically modern H. sapiens, there is

222

ET AL.

J. L. ARSUAGA

a midsagittal glabellar depression and/or the supraglabellar sulcus or fossa extends downwards in the glabellar segment, separating partial or totally one superciliary arch from its complement

on the opposite side. On the contrary,

zone appears continuous the browridge. “guessed”

in Neandertals

and AT-200

as a result of the complete lack ofa furrow separating

According

to Spitery ( 1985)) the Neandertal

to be present in Steinheim

and Bilzingsleben.

morphology

Actually,

the glabellar the two sides of

(his type A) can be

in Steinheim

the supra-

glabellar sulcus extends well inferiorly and the supercilliary archs are distinctly separated, but in Bilzingsleben there is only a slight indication ofglabellar depression (VlEek, 1978) and the torus appears continuous in the glabellar region. After examination of a cast, and from VlEek’s descriptions and figures, we conclude that the Bilzingsleben frontal bone shows a morphology similar to AT-200 and thus possesses derived Neandertal features (and is not, contra VlEek, 1978, Homo erectus-like). A Neandertal

derived character

is the circular

or oval cranial outline in norma occipitalir

(Arsuaga et al., 1989). This trait is already found in the European clearly in Biache 1). In the Atapuerca observe

the cranial

outline.

However,

large parietal bone fragments, struction of both biparietal

middle Pleistocene

(very

hypodigm no specimen is complete enough to directly Parietal

I and Parietal

II are two complementary

with a broad overlap region which permits a tentative recon-

vault morphologies.

The projected

outlines of the parietal walls

would be parallel or even slightly converging toward the top. We consider plesiomorphic with respect to the Neandertal apomorphic circular profile. Only the best preserved occipital

bone in the Atapuerca’s

hypodigm

(0.

this pattern

III)

permits a

true phylogenetic discussion. According to Hublin (1984), the Neandertal occipital bone displays an occipital torus showing a bilateral maximum development and a clearly defined suprainiac fossa on a strongly convex occipital plane (occipital protrusion). To Hublin ( 1982, 1984) these three Neandertal features are found associated in the Early Wurm (“classic”) Neandertals Delaunay,

as well as in the Riss-Wiirm Saccopastore

Neandertals

1 and 2, EhringsdorfS)

(Krapina,

La Chaise

and in some Riss specimens

Bourgeois-

(Biache and La

Chaise Suard 2). This author also recognizes the bilateral development of the torus and a faint and not well delimited suprainiac fossa in Steinheim and Swanscombe. On the contrary, the Atapuerca Occipital III does not exhibit the Neandertal derived morphology. There is no suprainiac fossa in 0. III and the fossil shows a rather flattened occipital plane compared with the “classic” Neandertals. In “classic” Neandertals, as a result of the strong convexity (protrusion) of the occipital plane, the opisthocranion is situated well above the nuchal ridge (and also above the suprainiac fossa). In 0. III the opisthocranion would be located in the nuchal ridge. According to Hublin ( 1982)) there is in Neandertals a depressed medial zone in the nuchal ridge between the two lateral protuberances. developed morphology features.

and not depressed of the occipiti

Size and morphology

near the midline.

Thus,

In 0. III the nuchal ridge is well 0. III

bone and not the characteristic

exhibits

the plesiomorphic

Neandertal

of the mastoid region have been commonly

association

of

used for phylogenetic

discussion. Some authors consider as a primitive feature the presence of a small mastoid process (Weidenreich, 1943; Endo & Kimura, 1970; Lumley & Sonakia, 1985; Hublin, 1986). However we are in agreement with Andrews ( 1984) in considering this character as not relevant for phylogenetic reconstruction in early and middle Pleistocene in Africa and Asia. Nevertheless, the lack of mastoid projection from the cranial base is generally judged to be a typical feature of NeandertaIs (Weidenreich, 1943; Patte, 1955; Vallois, 1969; Endo & Kimura, 1970; Santa Luca, 1978; Smith, 1980; Wolpoff, 1980a; Heim, 1981-82; Trinkaus,

CRANIAL

REMAINS

AND

LONG

BONES

FROM

ATAPUERCAIIBEAS

223

1983; Stringer et al., 1984; Tillier, 1986; Condemi, 1987; Mallegni & Radmilli, 1988). Some authors (Heim, 1974; Smith, 1980; Trinkaus, 1983) argue that the lack of mastoid projection from the cranial basis in Neandertals is not related to decrease of the mastoid process but to the inflation (pneumatization) of the cranial basis at the mastoid region. As a result of this extensive pneumatization the mastoid process appears enclosed in the petrous region and there is an evident increase in the caudal projection of the cranial base, specially the juxtamastoid eminence, which projects downward to the mastoid process. The presence of an eminence (or occipitomastoid crest) projected when compared with the mastoid process is judged by several authors as a Neandertal autapomorphy (Santa Luca, 1978; Stringer et al., 1984; Tillier, 1983, 1986; among others). In AT-84 and AT-86 the mastoid processes are well isolated from the petrous region and they are very projected from the cranial base (see Table 4). AT-84 preserves also the juxtamastoid eminence, which projects downward less clearly than the mastoid process. Therefore both Atapuerca specimens lack the mastoid pneumatization we consider autapomorphic for Neandertals. The mastoid process size has been used to determine the sex of middle Pleistocene fossils (Wolpoff, 1980a,6), Krapina (Smith, 1980) and Shanidar specimens (Trinkaus, 1983). On the other hand, Heim (1974, 1981-82) rejects the use of this trait in sex determination of “classic” Neandertals. Length and projection of mastoid process is represented in Figure 15. It can be seen that the positions in the scatter diagram of the Krapina temporal bones are consistent with the classification of males and females given by Smith (1980). Western Neandertals do not show such a clear dimorphism (in agreement with Heim’s assertions). The Atapuerca temporal bones show a disparity in mastoid process size which is consistent with the sexual dimorphism pattern ofthe Mediaeval Seplilveda sample. So, AT-84 would be a male and AT-86 a female. On the other hand, ifthe mastoid process projection is taken from the parietal notch the scatter diagram provides a more secure sex determination for the Sepulvedasample (Figure 16). Again, AT-84 must be considered a male and AT-86 (Cranium 1) a female. Furthermore, AT-84 shows more developed mastoid and supramastoid crests than AT-86. In AT-86 the tympanic plate is thick and can be divided into two parts. The anterior halfis scarcely convex and the posterior half is quite thick. These traits are present in Homo erectus (Weidenreich, 1943; Rightmire, 1984; Stringer, 1984) and Neandertals (Endo & Kimura, 1970; Tillier, 1984, 1986) and can be considered plesiomorphies. An external auditory meatus positioned superiorly has been considered a typical feature (or autapomorphic) of Neandertals (Patte, 1955; Vandermeersch, 1978; Tillier, 1983; Stringer et al., 1984). This position results in a location of the porus at the level of the zygomatic process. Also, the superior margin of the meatus lies substantially above the roof of the mandibular fossa. In ER-3883, ER-3733,OH-9 and Steinheim (observed from casts) the acoustic porus lies below the zygomatic level and its superior margin is at the level or slightly above the roof of the mandibular fossa. For us, this is the plesiomorphic state. In AT-86 the acoustic porus is below the zygomatic process and in AT- 124 the superior edge of the porus is at level of the mandibular fossa roof. So, both Atapuerca fossils exhibit the plesiomorphic condition. A wide and shallow mandibular fossa is mentioned as a typical (or autapomorphic) attribute of Neandertals (Vallois, 1969; Heim, 1974; Vandermeersch, 1978; Tillier, 1983, 1986). On the other hand, a mandibular fossa deep and narrow in anteroposterior direction can be found in “Sinantho~us” (Weidenreich, 1943)) Caste1 di Guido (Mallegni & Radmilli,

J. L. ARSUAGA ET AL.

* * *

sr

+ +

*

*

lk *

*

**

*

* ::

*

sr*

a**

* sr

*

‘0””

0

**

kLQ5 **

kAT-&

#

“gy2

&I

@

LQ%J L 27 8 &jLFl

*

OW. E NEANDERTALS @ATAPUERCA/

20

22

24

MASTOID

PROCESS

26

EASE

28

30

IsEAs

32

34

LENGTH

Figure 15. Scatterdiagram for length of the mastoid process base and projection of the mastoid process (Zoja’s method). LFl = La Ferrassie 1; LCH = La Chapelle-aux-Saints; GBI = Gibraltar I; LQ5 = La Quina 5; LQlO = La Quina 10; LQ27 = La Quina 27.

1988),

ER-3733,

ER-3883,

OH-9

and Steinheim

(on casts). The last condition

must be

considered plesiomorphic and AT-124 shows it. To some authors (Vallois, 1969; Endo & Kimura, 1970; Heim, 1974; Vandermeersch, 1978) a well developed postglenoid process is a typical Neandertal feature. In Tillier’s opinion ( 1983, 1986) prominent postglenoid uncertain. A phylogenetic traits are metrical

this is a plesiomorphy for Neandertals. AT- 124 displays a very process but for us the phylogenetic status of the character remains

analysis of postcranial (continuous)

characters

and so they cannot

is very difficult at present because most be easily broken down into character

Figure 16. Scatterdiagram

**a

Sr+r

PROJECTION

34

*

*

*

36

*

*

FROM

*

*

*

r.c

38

*

*

**

2r

*

*

sr sr

sr

*

4

*

40

a

6

*

AT-86

+

*

sr

for length ofthe mastoid process (Zoja’s method)

32

PROCESS

*

30

*

*

MASTOID

i?

*

*

*

*

*

t

44

*

*

Q

+*

*

*

46

It

t

**

*

*

*

**

*

t

*

48

*

50

*

*

52

*

54

(I.P.) to the tip ofthe mastoid process.

**

*

84

BD, AT-

and distance from the incisuraparictalis

42

*

*6 *

t

#

t

*

*

1

226 states

J. L. ARSUAGA

(discrete).

Moreover

the ranges

ET AL.

of most metrical

variables

show ample

overlap

between modern and fossil human samples. Finally, the postcranial remains are scarce (or absent for many anatomical regions) in the European middle Pleistocene record. In spite of this, in the following discussion we will point out the features that the Atapuerca with Neandertals

or modern populations.

The well developed hypertrophied

fossils share

lesser tubercle

of AT-25

(and probably

AT-93)

corresponds

to a

m. subscupuluris. There is also in both fossils a deep fossa for the gleno-humeral

inferior ligament. These two traits suggest a musculo-ligamentous hypertrophy related to the maintenance of the articular integrity. The great cortical thickness of the three Atapuerca humeri

(compared

with modern

humeri)

would also reflect upper limb robusticity.

This

biomechanical pattern has been claimed for Neandertals by several authors (Trinkaus, 1977, 1982, 1983, 1984, 1989; Endo & Kimura, 1970; Thoma, 1975; Trinkaus & Howells, 1979; Smith, 1984; Ben-Itzhak et al., 1988). On the other hand, the internal surface of AT-25 and AT-93 is flat and the deltoid tuberosity broad, as in modern humans. In Neandertals the internal

surface is typically

morphology

shared by the Atapuerca

plesiomorphic diameter

convex and the deltoid tuberosity

state and that of Neandertals

of humeral

narrow.

the apomorphic

state. In AT-25

head is equal or greater than the vertical diameter.

lations the vertical diameter dence of this anatomical

Therefore,

the

humeri and modern humans seems to represent

is always the greatest.

Unfortunately,

the

the transverse

In modern popu-

because the scanty evi-

region in the fossil record the phylogenetic

status of this trait is

uncertain. Since AT-25 and AT-93 correspond to the same side (right) and are probably of similar age, the strong disparity between them in development of the muscular attachments can be due either to sexual dimorphism or to handedness. In the Atapuerca/Ibeas sample handedness has been established for 23 anterior teeth belonging to at least eight individuals and all of them were found to be right-handed consider sexual dimorphism The most remarkable amygdaloid thickness.

de Castro, personal communication),

so we

feature of the Atapuerca

tibiae is robusticity,

which is reflected in a

cross section shape at both midshaft and 2/3 levels and also in a great cortical The Atapuerca

populations. following

(Bermudez

as the most plausible explanation.

Furthermore,

traits:

tibiae are in these traits closer to Neandertals the specimens from Atapuerca

weakly developed

shallow fibular articular Pleistocene fossil record,

anatomical

margins,

than to modern

share with other Neandertals flat distal articular

surface

the and

surface. Since there is no other tibiae in the European middle it is not possible to ascertain whether these character states are

plesiomorphies or apomorphies. In sum, the Atapuerca/Ibeas

cranial

and postcranial

sample shows a number

of plesio-

morphic traits (i.e., character states not retained by the upper Pleistocene Neandertals but present in European middle Pleistocene fossils as well as in the early or middle Pleistocene out of Europe). Also some Neandertal derived features have been recognized (i.e., character states never found out of the Neandertal geographic area). Furthermore, plesiomorphies and Neandertal

apomorphies

1989). The Wi.irm Neandertals

are found in the Atapuerca/Ibea.s form an homogeneous

mandibles sample (Aguirre et al.,

group, with a very characteristic

pattern of

apomorphic features despite some differences among temporal or geographically distant samples. The Riss-Wiirm European samples must also be considered fully Neandertal with most (or even all) the Neandertal apomorphies. In the European middle Pleistocene BiacheSaint-Vaast 1 (Riss) and the parietal vault, occipital and temporal bones from La Chaise-

CRANIAL

REMAINS

AND

LONG

Suard (final Riss) appear fully Neandertal Other middle Pleistocene

crania

BONES

FROM

regarding the characters

(Steinheim,

227

ATAPUERCA/IBEAS

Swanscombe,

of the preserved regions.

Arago and Petralona

most complete) have been claimed to present some Neandertal definitely not Neandertals because of the overall plesiomorphic

are the

apomorphies, but they are morphology. Furthermore

the derived features are not shared by all the European middle Pleistocene fossils but vary from one another. The problem with classifying these European middle Pleistocene fossils is that they cannot be defined in a strictly cladistic manner, simply because they do not possess uniquely derived character states. The plesiomorphic traits they exhibit are shared with nonEuropean contemporaries or ancestors, and their derived features (when present) are shared with Neandertals. This is a problem of all paraphyletic groups (in Hennigian systematics those groups that have a common ancestry but from which the descendant groups have been excluded).

So, we have to define them in terms of the absence of the distinctive characters of

their descendants

and from the presence of characters

absent in their ancestors

(Carroll,

1988).

Acknowledgements Thanks

to Dr J.

supported

Azpeitia

by the Direction

for the radiographies General

he made for us. This study has been

de Investigation

Cientifica

y TCcnica,

Project

No.

PB86-06 15403-02.

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ATAPlJERCA/IBEAS

229

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Resumen En el presente

trabajo

se analizan

las relaciones

Huesos de Cueva Mayor (Sierra de Atapuerca, evolution humana en el Pleistocene y variables metricas de1 crineo tica.

Los

estados

Neandertales

de 10s fosiles de la Sima de 10s Burgos) en el context0 de la

europeo. El estudio se refiere a 10s caracteres

morfologicos

y huesos largos, que se abordan desde una perspectiva

de 10s caracteres

de1 Pleistocene

filogeneticas

Ibeas de Juarros,

reflejan:

superior

(a)

pero presentes

plesiomorhas

no

retenidas

en fosiles de1 Pleistocene

cladispor

10s

medio

e

inferior (en Europa y fuera de ella) asignados a Homo erectus y Homo sapiens; (b) una apomorfia compartida con 10s Neandertales; (c) algunos caracteres postcraneales compartidos con 10s Neandertales cuyo significado filogenetico no ha sido establecido. En consecuencia, 10s fosiles de Ibeas/Atapuerca estan filogentticamente relacionados con 10s Neandertales, pero no pueden ser considerados Pleistocene medio Neandertales

el mismo grupo a causa de las plesiomorfias retenidas. Otros fosiles de1 europeo muestran igualmente apomorfias compartidas con 10s

(aunque

no necesariamente

las mismas en todos 10s cases), junto

morfias. En nuestra opinion 10s fosiles de1 Pleistocene derivados

exclusives

y por tanto

no pueden

ser agrupados

TambiCn se estudian otros aspectos coma el dimorfismo coma la lateralizacion

de1 cerebra.

con plesio-

medio europeo no comparten y definidos

rasgos

cladisticamente.

sexual en el humero y temporal,

asi